BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, researchlibraries, and research funders in the common goal of maximizing access to critical research.

Rediscovering Dyckia excelsa (Bromeliaceae) in Mato Grosso do Sul, Brazil:Taxonomy, Geographic Distribution, and Notes on Leaf AnatomySource: Systematic Botany, 40(1):129-135.Published By: The American Society of Plant TaxonomistsURL: http://www.bioone.org/doi/full/10.1600/036364415X686422

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, andenvironmental sciences. BioOne provides a sustainable online platform for over 170 journals and books publishedby nonprofit societies, associations, museums, institutions, and presses.

Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance ofBioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiriesor rights and permissions requests should be directed to the individual publisher as copyright holder.

Systematic Botany (2015), 40(1): pp. 129–135© Copyright 2015 by the American Society of Plant TaxonomistsDOI 10.1600/036364415X686422Date of publication February 12, 2015

Rediscovering Dyckia excelsa (Bromeliaceae) in Mato Grosso do Sul, Brazil:Taxonomy, Geographic Distribution, and Notes on Leaf Anatomy

Gecele M. Paggi,1,4 Rafael B. Louzada,2 Iria H. Ishii,1 Adriana Takahasi,1

Rosani C. O. Arruda,3 and Aline P. Lorenz-Lemke3

1Ciencias Biologicas, Campus do Pantanal, Universidade Federal de Mato Grosso do Sul, P. O. Box 252,Corumba, Mato Grosso do Sul, 79304-902 Brazil.

2Centro de Ciencias Biologicas, Departamento de Botanica, Universidade Federal de Pernambuco,Recife, Pernambuco, 50940-000 Brazil.

3Centro de Ciencias Biologicas e da Saude, Universidade Federal de Mato Grosso do Sul,Campo Grande, Mato Grosso do Sul, 79070-900 Brazil.4Author for correspondence (gecelepaggi@gmail.com)

Communicating Editor: Jimmy Triplett

Abstract—In this contribution the rediscovery of the bromeliad Dyckia excelsa in Mato Grosso do Sul state, central-western Brazil, isreported. This species was first described based on a single individual from a particular collection in 1993 with no precise locality.Illustrations and a brief discussion on the leaf anatomy of the species are provided. One additional species, Dyckia gracilis, which has beenoverlooked by previous authors, is included in the list of Bromeliaceae of Mato Grosso do Sul State. An updated key for Dyckia species fromthe region of D. excelsa is presented. The status of D. excelsa in Mato Grosso do Sul State, Brazil, is discussed, and recommendations for itsconservation are offered.

Keywords—Estrada Parque Pantanal, Pantanal wetlands, Pitcairnioideae, rock outcrops, South America.

The phytogeography of the Mato Grosso do Sul State (MS),in central-western Brazil, comprises 50% of wide-rangingspecies, 30% of Cerrado species, and 20% of species sharedamong other origins (Pott and Ratter 2011). Those species aredistributed in three different biomes: Cerrado (Savanna), thePantanal wetland (including part of the Chaco), and part ofthe Atlantic Forest (IBGE 2004; Junk et al. 2011). The Pantanalbiome is severely threatened, and it is considered one ofthe areas of heaviest human impact in MS (Cavalcanti andJoly 2002; Junk et al. 2011). The ironstone outcrops called“bancadas laterıticas” or “canga” can be considered a pecu-liar feature of the Central Brazilian landscape. They are clas-sified in the Humid Area as intermittent terrestrial systems,being rocks covered periodically with water for varyinglengths of time, with or without sparse vegetation (Nunes daCunha and Junk 2011; Junk et al. 2014).

Dyckia Schult. & Schult. f. (1830: 1194) is the second largestgenus in the Pitcairnioideae (Bromeliaceae) with about 160species (Butcher and Gouda 2013) distributed in southern,southeastern, western and northeastern Brazil, northernArgentina, northern Paraguay and northern Uruguay (Smithand Downs 1974; Givnish et al. 2007, 2011, 2014). In Brazil,it is represented by around 130 species of which 112 areendemic (Forzza et al. 2014). Dyckia species can be found assingle segregated individual plants established by dispersedseeds or forming large and dense clusters. They are usuallyxeromorphic terrestrials growing in well-drained soils, orrupicolous on diverse rock outcrops (i.e. granitic, quartzitic,sandstone, or iron-rich “canga”) of dry or sometimes humidhabitats (Smith and Downs 1974; Strehl 1994; Wiesbauer et al.2007). Recently, some species of Dyckia genus were describedand identified as endemic species of rock outcrops from MS(Versieux et al. in press): D. pauscispina Leme & E. Esteves(Leme and Pereira 2003), D. grandidentata P. J. Braun &Esteves (Braun and Pereira 2008), D. stolonifera P. J. Braun &Esteves (Braun and Pereira 2009) and D. pottiorum Leme(Leme et al. 2012), showing the great importance of theregion for conservation issues.

After examining some particular specimens from a naturalpopulation of the genus Dyckia collected during recent fieldexpeditions in the Estrada Parque Pantanal, close to the bor-der of Brazil with Bolivia, we concluded that those specimensbelong to Dyckia excelsa Leme (1993: 6). This species wasdescribed in 1993 based on a collection of Roberto Burle Marx(R. Burle Marx s. n.), which flowered in cultivation. Sincethen, no record or information about this species has beenfound in herbaria or the literature to elucidate the exact local-ity where this taxon occurs.Dyckia excelsa corresponds to the eleventh species of Dyckia

recorded for MS, including one other species (Dyckia gracilisMez) which has been cited in the literature (Smith andDowns 1974) but was missing in the Brazilian Bromeliaceaechecklist (Forzza et al. 2014). Thus, herein we provideupdated information about taxonomy, geographic distribu-tion, and leaf anatomy as well as a key for the species ofDyckia from the Pantanal region, aiming to contribute to thefuture monograph of Bromeliaceae for the Flora of the Stateof Mato Grosso do Sul.

Material and Methods

The morphological description of the species is based on measure-ments of living plants taken during field work in the ironstone outcropsclose to the Pantanal of Mato Grosso do Sul State, Brazil, and examinationof herbarium specimens inspected at the COR (UFMS/CPAN) and CPAP(EMBRAPA Pantanal) herbaria listed in the Results section. Field obser-vations of the populations of this species were conducted by I. H. Ishiiand G. M. Paggi during their fieldwork in the Estrada Parque Pantanaland BR-262, Corumba city, from 2009–2013. Conservation assessment forthe species was based on field observation, applying the IUCN red listcategory criteria (IUCN 2012). Information about flower, fruit, and seedcolor was taken mainly from our own observations in the field. Thegeographic distribution for each species is based on fieldwork at naturalpopulations. For leaf anatomy, light microscopy observations were per-formed using mature leaves fixed in 30–40% formalin: acetic acid: 50%ethanol (FAA, 18:1:1), dehydrated in ethanol series, and stored in 70%ethanol (Johansen 1940). Anatomical preparations were taken midwaybetween the apex and base of the lamina. Leaf segments were embeddedin polyethylene glycol 1500 (PEG) without softening (Barbosa et al. 2010).

129

The sections were stained with 2% alcian blue and 0.1% basic fucsin in50% ethanol (Luque et al. 1996).

Results

Dyckia excelsa Leme (1993: 6).TYPE: BRAZIL. Mato Grosso,sem local preciso. November 1992 (fl. in cult.) R. BurleMarx s. n. (holotype: HB!; isotype: RB!).

Saxicolous, short to long caulescent, propagating by basalshoots, 70–130 cm high. Leaves arranged in a rosette; leafsheaths, ca. 7 + 6 cm, coriaceous, transversal elliptic, bright-castaneous, glabrous on both surfaces, margins serrulate atthe apex, entire toward the base; leaf blades ca. 58 + 3 cm,stiff coriaceous, flat at the base, canaliculate toward the apex,linear-triangular, green, abaxial surfaces lepidote, adaxial sur-faces glabrous, scales white, adpressed, margins lax-serrate,prickles ca. 5 mm long, ca. 7 mm apart, triangular, complanate,green to castaneous, antrorse or straight at the base, apexpungent. Inflorescences 70–130 cm long; peduncles 40–100 cmlong, terete, axyllary, densely lepidote, scales ferrugineous atthe base, green; peduncle bracts 0.5–1.2 +3.7–10.2 cm, shorterto longer than internodes, coriaceous, erect, green, glabrous,margins serrate, apex acuminate; fertile part of the inflores-cence in racemes, ca. 30 cm long, linear, rachis conspicu-ous, erect, woolly-lepidote, green. Floral bracts 0.3–0.9 +0.5–3.6 cm, coriaceous, triangular, ecarinate, margins entire,castaneous, sparsely woolly-lepidote on both surfaces, apexacute. Flowers odorless ca. 40 mm long (excluding pedicel),tubular; pedicel ca. 10 mm long; sepals ca. 15 + 7 mm, ellip-tic, abaxial surface woolly-lepidote, scales subadpressed,orange, symmetric, ecarinate, apex emarginate; petals 24–29 + 10–12 mm, elliptic, abaxial surface lepidote, adaxialsurface glabrous, margins entire, orange, winged, wingswhite (in sicco), apex obtuse; filaments 30 mm long, free,complanate, white; anthers 11 mm long, subbasifixed, basesagittate, yellow, apex acute, recurved at the anthesis. Anthe-sis occurs through 12 h, with the first flowers starting early inthe morning, and three to six flowers opening subsequently.Ovary ca. 15 mm long, obclavate, green; placentation axial.Stigma conduplicate-spiral, yellow. Fruit a capsule, widelyovate. Seeds 5 + 2.5 mm, flat, asymmetric, red to brown, wingspale brown. Figures 1A–M and 2 A–I.Etymology—The name Dyckia excelsa refers to the tall habit

that confers a special aspect to the plant.Phenology—Flowering from April to November and

fruiting from May to January. Each rosette can produce upto three inflorescences at the same time.Distribution and Habitat—So far the species is only

known from Corumba and Ladario cities, in the westernportion of the state of Mato Grosso do Sul, central-westernBrazil, restricted to rock outcrops (granitic and iron-rich),next to the highlands in the Residual Plateau of Urucum, onthe border of Brazil and Bolivia (Figs. 2B and 3).Ecology—It grows forming dense heliophylous popula-

tions on patches of exposed calcareous rock and ironstoneoutcrops, deciduous forest, inside Pantanal Bioma. OtherBromeliaceae species that occur in the same region includeAnanas ananassoides (Baker) L. B. Smith, Bromelia balansae Mez,Deuterocohnia meziana Kuntze ex Mez, Dyckia leptostachyaBaker, Dyckia gracilis Mez, Tillandsia loliacea Martius exSchultes f., T. streptocarpa Baker, and T. duratii Visiani.Conservation—The species is treated as endangered (EN),

due to its occurrence in only one large population (Fig. 2B) at

the Farm of Sao Joao and three more little populations in theCorumba and Ladario cities (BR-262 and Marina Gatass Park),none of which is protected within Brazilian national parks.

Comparison—According to the diagnosis of Dyckia excelsa(Leme 1993), this species is morphologically related to Dyckiaferruginea Mez, differing mainly by flowers about 5 cm longsupported by a pedicel 6–10 mm long, sepals twice as bigand petals three times as long and orange.

Based on the key of Smith and Downs (1974), Dyckia excelsais also morphologically related to D. insignis Hassl. (1919:316) due to the long exserted stamens. Although they are notsympatric species, both occur with certain proximity innorthern Paraguay and the northern part of the State of MatoGrosso do Sul in Brazil. After the analysis of morphologicalcharacters, we conclude that they differ one from anothermainly because D. excelsa is taller when flowering (70–130 cm high vs. 30–40 cm high), with longer leaves (ca. 65 cmvs. 20–35 cm), bigger flowers (ca. 40 mm long vs. 22–28 mmlong) and the corolla almost closed at anthesis (vs. openedat the anthesis). Besides these, two other species (D. gracilisand D. leptostachya Baker) are found in the same region asD. excelsa. The main morphological differences among themare summarized in Table 1.

Additional specimens examined—BRAZIL. Mato Grosso do Sul: RodoviaRamon Gomez, Parque Marina Gattass, 19�00047.300S, 57�41018.400W, 100 melev., 07 Nov 2004, M.C. Mendes et al. 4 (COR); ibidem, Fazenda Sao Joao,Estrada Parque Pantanal, MS-238, 19�1007700S, 57�32031.800W, 88 m elev., 29Nov 2006, A. Takahasi et al. 1098 (COR); ibidem, Ladario, V. J. Pott et al. 208(CPAP); ibidem, 19�1007700 S, 57�32031.800 W, 88 m, 31 October 2012, I. H.Ishii et al. 785 (COR, UFP).

Leaf Anatomy—The leaf epidermis is composed of com-mon epidermal cells, stomata, and multicellular peltate tri-chomes (scales). In front view, the common epidermal cellsare oblong-shaped, and have sinuous, lignified, and thickwalls. A spherical silica body is observed in each cell. Onboth surfaces, the peltate trichomes are composed by roundedcells, nearly concentrically organized with conspicuously thinprimarywalls, inflated in appearance. The trichomes are shield-like, sparsely distributed on the adaxial surface, and numerouson the opposite surface where the stomata are located. On thelower surface, the epidermis is organized in costal and inter-costal bands or zones (Figs. 4A and B). In the costal zones, onlycommon epidermal cells are present and they have stronglythick cell walls. Stomata and trichomes are restricted to epider-mal furrows and the trichomes shield forming an indumentumthat overlaps all cells.

In cross section, both surfaces are strongly ribbed on theunderside of the leaf blade (Figs. 4B and E). The epidermis isone-layered and the common epidermal cells are flattenedand exhibit thick internal cell walls (Figs. 4B, E, and H). Thestomata, restricted to the lower surface, are located ingrooves and covered by a shield of peltate trichomes(Figs. 4B, I, and J). A hypoderm with strongly thickened cellwalls and with no chloroplasts occurs under the upper epi-dermis forming two or three layers (Figs. 4H). The hypoder-mal cells are septate fiber-like and lignified. In lower surface,the 4–5 hypodermal layers produce ribs under epidermalcostal. These projections produce grooves where the stomataare located.

Under the upper hypodermis we observed long, thinwalled and voluminous cells similar to palisade parenchyma,occupying almost 3/4 of the upper mesophyll (Fig. 4A). Smallchloroplasts are observed in this tissue (Figs. 4C and D). Theremaining mesophyll is filled with rounded or short-armed

130 SYSTEMATIC BOTANY [Volume 40

Fig. 1. A–M. Dyckia excelsa. A. Habit. B. Detail of the inflorescence apex. C. Detail of the leaf sheath. D. Detail of the leaf blade. E. Detail of theprickle. F. Flower. G. Flower without calix and corolla. H. Petal. I. Sepal. J. Detail of the anther. K. Detail of the stigma. L. Fruit immature. M. Fruit mature.

2015] PAGGI ET AL.: REDISCOVERING DYCKIA EXCELSA 131

Fig. 2. A–I. Dyckia excelsa, Fazenda Sao Joao, Estrada Parque Pantanal, Corumba, Mato Grosso do Sul. A. Habit of adult individual in naturalpopulation. B. General aspect of an ironstone outcrop, showing a population ofD. excelsa in dry vegetation and soil, with the Residual Plateau of Urucumon the back. C. Fertile individuals of D. excelsa growing near the typical vegetation. D. Juvenile individuals of D. excelsa growing associated withDiscocactus ferricola Buining & Brederoo. E. Detail of dehiscent fruit with seeds inside. F. Detail of flower at the anthesis; note the densely lepidoteindumentum on petals and sepals and the stamens’ curvature. G. Detail of conspicuous rhizome characteristic of this species. H. Detail of the abaxialsurface of the leaf blade, showing the scales arranged in rows along the midnerves and brown prickles arranged in different directions. I. Variation inpeduncle height found in the natural population. (Photos: I. H. Ishii: A, B, C, and G; G. M. Paggi: D, E, F, H, and I).

132 SYSTEMATIC BOTANY [Volume 40

chlorophyllous cells (Figs. 4D and 4F). The arm cells delimitlarge air spaces; they occur in the lower mesophyll connectingchlorenchyma with the sub-stomatal chamber. Idioblasts withraphide crystals are restricted to near the vascular systembelow long parenchyma cells (Figs. 4D and G). The vascularsystem of the leaf blade is composed of collateral vascularbundles. A prominent parenchyma sheath, an endodermis, isidentified. Inner parenchyma sheath lignified fibers surroundthe vascular tissue completely (the large bundles) or partially.i.e. surrounding only the phloem region, in the small-diametervascular bundles (Figs. 4D and E).

The anatomical features of the leaf of this species confirmits position in the “xeric clade” (Givnish et al. 2007; Santos-Silva et al. 2013), considering the plant habitat environmentalconditions most of the year: seasonally dry tropical forests,with summer temperatures reaching 49�C and with severe

winter frosts (Pennington et al. 2000). Attributes observed inthe epidermis, such as the sinuous shape of ordinary epider-mal cells, protected stomata restricted to intercostal bands onthe abaxial surface, and the hypodermis differentiated intoperipheral rigid mechanical and internal water-tissue of sev-eral layers, are included as key innovations and may haveallowed bromeliads to invade new adaptive zones in aridregions (Givnish et al. 2014). The species grows in rocky soilwith a minimum amount of substrate, which makes impor-tant the accumulation of water and nutrients in the leaves.The presence of long cells with chloroplasts in the hypoder-mis can make the photosynthetic process more efficient: thickleaf blades are consistent with a non-tank growth formand CAM photosynthesis of the xeric clade (Santos-Silvaet al. 2013; Givnish et al. 2014). Here we identified most ofdescriptions by Tomlinson (1969). The organization of multi-cellular peltate trichomes, with no radial pattern, confirmsthe observations of Varadarajan and Gilmartin (1987) forPitcairnioideae. However, internal hypodermic layers differ-entiated into chlorenchymatous water-tissue are a distinctivecharacter of this species now described but not previouslymentioned for Dyckia, which is frequently described as hav-ing water-tissue of colorless cells.Final remarks—Dyckia comprises more than 160 terrestrial

or epilithic species with robustly xeromorphic habits. Most ofits members belong to the azonal rock vegetation of Neotrop-ical savannas and forests of Brazil and adjacent countries(Smith and Downs 1974). High levels of species richness anda large proportion of narrow endemics of this genus havebeen described in “rocky fields” (Versieux and Wendt 2006,2007; Versieux et al. 2008). We found D. excelsa occurring onlyin rock outcrops of MS and its occurrence inMato Grosso state(MT) should be taken with caution, since no collections of thisspecies were found in the herbaria research.The total number of Bromeliaceae species from MS was

described as 43, and at least five endemic species (Versieuxet al. in press), four Dyckia species (Braun and Pereira 2008,2009; Leme et al. 2012) and Tillandsia bonita Versieux &Martinelli (Versieux et al. 2013), all of them suffering fromhabitat loss. Based on data collected for this study (see alsoSpeciesLink, www.splink.org.br), we hypothesized that D.excelsa may be also endemic in MS. This study describeddifferent aspects of the biology of D. excelsa, adding anatomicand geographic information for this taxon. Accurate knowl-edge of taxonomy and geographic distribution of species is ofgreat importance for phylogeny and phylogeography studiesthat have been trying to explain the radiation and evolutionin this recent group (Krapp et al. 2014).

Artificial Key to the Species of DYCKIA from Pantanal Region, Mato Grosso do Sul, Brazil

1. Flowers ca. 40 mm long, long pedicelate; sepals obtuse; corolla tubular; stamens not connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. excelsa1. Flowers 13–23 mm long, subsessile; sepals emarginate; corolla infundibuliform; stamens connate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. Leaves 40–100 cm long; flowers orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. leptostachya2. Leaves ca. 30 cm long; flowers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. gracilis

Acknowledgments. We would like to thank Marcus VinıciusSantiago Urquiza, Brenda Baıa Brandao, Diego Finati Alves, FlorenceCristina dos Santos Ignacio, Gilaine Moreira de Miranda, Kelly ConceicaoRondon de Arruda, Letıcia Helena de Moraes Delmao, and Maria ReginaPimentel Montano for help in field and lab work. Thanks to Mr. Diogo

Silva Matos for assistance with anatomical slide preparations and ReginaCarvalho for the line drawings. RCOA gratefully acknowledges CNPq fora research grant. This work was supported by grants from FUNDECT(Fundacao de Apoio ao Desenvolvimento do Ensino, Ciencia e Tecnologiado Estado de Mato Grosso do Sul) to G. M. Paggi.

Fig. 3. Map of geographical distribution of Dyckia excelsa in theBrazilian Pantanal, showing all four populations observed in Corumbaand Ladario cities in the state of Mato Grosso do Sul, Brazil, in the borderwith Bolivia.

Table 1. Comparison among Dyckia excelsa and the two other knownspecies occurring in Corumba and Ladario cities, MS.

D. excelsa D. leptostachya D. gracilis

Leaves (cm) ca. 65 40–100 30Prickles (mm) ca. 5 3–4.5 ca. 4Pedicel Long Short ShortFlowers (mm) ca. 40 13–23 ca. 18Corolla Closed Opened OpenedSepal apex Obtuse Emarginate EmarginatePetal color Orange Orange Yellow

2015] PAGGI ET AL.: REDISCOVERING DYCKIA EXCELSA 133

Fig. 4. A–J. Leaf anatomical characteristics of Dyckia excelsa. A–F, H–J: cross sections. G: paradermal section. A. Leaf margin. B. Detail of epidermisand hypodermis. C. Detail of long water hypodermal cells with chloroplasts (arrows). D. General aspect of mesophyll showing chlorenchyma cells,raphide-sacs (empty, see arrows). E. Details of inner part of mesophyll: chlorenchyma, and sclerotic layers of hypodermis, epidermis and collateralvascular bundles (arrows). F. Detail of arm-cells and minute intercellular spaces (ac). G. Raphide-sacs. H–J. Details of epidermis and hypodermal cells:reduced lumen, thick cell walls, stoma and peltate trichome.

134 SYSTEMATIC BOTANY [Volume 40

Literature Cited

Barbosa, A. C. F., M. R. Pace, L. Witovisk, and V. Angyalossy. 2010. Anewmethod to obtain good anatomical slides of heterogeneous plantparts. IAWA Journal 31: 373–383.

Braun, P. J. and E. E. Pereira. 2008. Succulent and Xeromorphic Bro-meliads of Brazil Part 4. Dyckia richardii and D. grandidentata, twonew succulent and heavily-armed species from west-central Brazil.Cactus and Succulent Journal 80(6): 319–324.

Braun, P. J. and E. E. Pereira. 2009. Succulent and Xeromorphic Brome-liads of Brazil Part 6. Dyckia stolonifera? A new and nearly extinctspecies from Mato Grosso do Sul. Cactus and Succulent Journal 81(6):301–304.

Butcher, D. and E. J. Gouda. 2013 (continuously updated). The new brome-liad taxon list. http://BromTaxonList.florapix.nl. Utrecht: UniversityBotanic Gardens (accessed 15–10–2013).

Cavalcanti, R. B. and C. A. Joly. 2002. Biodiversity and conservationpriorities in the cerrado region. Pp. 351–367 in The cerrados of Brazil,ecology and natural history of a neotropical savanna, eds. P. S. Oliveiraand R. J. Marquis. New York: Columbia University Press.

Forzza, R. C., A. Costa, J. A. Siqueira-Filho, G. Martinelli, R. F. Monteiro,F. Santos-Silva, D. P. Saraiva, B. Paixao-Souza, R. B. Louzada, andL. M. Versieux. 2014. Bromeliaceae in Lista de especies da Flora doBrasil. Jardim Botanico do Rio de Janeiro. Available from: http://reflora.jbrj.gov.br/jabot/floradobrasil (accessed: 02 May 2014).

Givnish, T. J., K. C. Milliam, P. E. Berry, and K. J. Sytsma. 2007. Phylogeny,adaptive radiation, and historical biogeography of Bromeliaceaeinferred from ndhF sequence data. Aliso 23: 3–26.

Givnish, T. J., M. H. Barfuss, B. Van Ee, R. Riina, K. Schulte, R. Horres,P. A. Gonsiska, R. S. Jabaily, D. M. Crayn, J. A. C. Smith, K. Winter,G. K. Brown, T.M. Evans, B. K. Holst, H. Luther, W. Till, G. Zizka, P. E.Berry, and K. J. Sytsma. 2011. Phylogeny, adaptive radiation, andhistorical biogeography in Bromeliaceae: Insights from an eight-locusplastid phylogeny. American Journal of Botany 98: 872–895.

Givnish, T. J., M. H. Barfuss, B. Van Ee, R. Riina, K. Schulte, R. Horres,P. A. Gonsiska, R. S. Jabaily, D. M. Crayn, J. A. C. Smith, K. Winter,G. K. Brown, T.M. Evans, B. K. Holst, H. Luther, W. Till, G. Zizka, P. E.Berry, and K. J. Sytsma. 2014. Adaptive radiation, correlated andcontingent evolution, and net species diversication in Bromeliaceae.Molecular Phylogenetics and Evolution 71: 55–78.

IBGE. 2004. Mapa Biomas do Brasil (1:5.000.000). http://mapas.mma.gov.br/geonetwork/srv/en/metadata.show?currTab=simple&id=298.Accessed 30 August 2013.

IUCN. 2012. IUCN red list categories and criteria: version 3.1. Gland,Switzerland and Cambridge, U. K.: IUCN Species Survival Commis-sion. 2nd Edition.

Johansen, D. A. 1940. Plant microtechnique, New York: Mc Graw-HillBook Co.

Junk, W. J., C. J. da Silva, C. Nunes da Cunha, and K. M. Watzen. 2011.The Pantanal: Ecology, biodiversity and sustainable management of a largeNeotropical seasonal wetland. Sofia, Moscow: Pensoft Publishers.

Junk, W. J., M. T. F. Piedade, R. Lourival, F. Wittmann, P. Kandus, L. D.Lacerda, R. L. Bozelli, F. A. Esteves, C. Nunes da Cunha, L. Maltchik,J. Schongart, Y. Schaeffer-Novelli, and A. A. Agostinho. 2014.Brazilian wetlands: their definition, delineation, and classificationfor research, sustainable management, and protection. Aquatic Con-servation: Marine and Freshwater Ecosystems 24: 5–22.

Krapp, F., D. S. B. Pinange, A. M. Benko-Iseppon, E. M. C. Leme, and K.Weising. 2014. Phylogeny and evolution of Dyckia (Bromeliaceae)

inferred from chloroplast and nuclear sequences. Plant Systematicsand Evolution, doi: 10.1007/s00606-014-0985-0.

Leme, E. M. C. 1993. Novas bromeliaceaes nativas do Brasil – XI. Pabstia4(4): 6–7.

Leme, E. M. C. and E. E. Pereira. 2003. New Bromelia and Dyckia species(Bromeliaceae) from Central Brazil. Vidalia 1(1): 21–30.

Leme, E. M. C., O. B. C. Ribeiro, and Z. J. G. Miranda. 2012. New speciesof Dyckia (Bromeliaceae) from Brazil. Phytotaxa 67: 9–37.

Luque, R., H. C. Sousa, and J. E. Kraus. 1996. Metodos de coloracao deRoeser (1972) – modificado e Kropp (1972) visando a substituicao doazul de astra por azul de alciao 8gs ou 8gx. Acta Botanica Brasilica10(2): 199–212.

Nunes da Cunha, C. and W. J. Junk. 2011. A preliminary classification ofhabitats of the Pantanal of Mato Grosso and Mato Grosso do Sul, andits relation to national and international wetland classification sys-tems. Pp. 127–141 in The Pantanal: Ecology, biodiversity and sustainablemanagement of a large neotropical seasonal wetland, eds. P Junk, W. J.,C. J. da Silva, C. Nunes da Cunha, and K. M. Watzen. Sofia, Moscow:Pensoft Publishers.

Pennington, R. T., D. E. Prado, and C. A. Pendry. 2000. Neotropicalseasonally dry forests and Quaternary vegetation changes. Journal ofBiogeography 27: 261–273.

Pott, A. and J. A. Ratter. 2011. Species diversity of terrestrial plants andhuman impact on the vegetation of the Pantanal. Pp. 281–300in The Pantanal: Ecology, biodiversity and sustainable managementof a large neotropical seasonal wetland, eds. P Junk, W. J., C. J.da Silva, C. Nunes da Cunha and K. M. Watzen. Sofia, Moscow:Pensoft Publishers.

Santos-Silva, F., D. P. Saraiva, R. F. Monteiro, P. Pitab, A. Mantovani,and R. C. Forzza. 2013. Invasion of the South American dry diagonal:What can the leaf anatomy of Pitcairnioideae (Bromeliaceae) tell usabout it? Flora 208: 508–521.

Smith, L. B. and R. J. Downs. 1974. Bromeliaceae (Pitcarnoideae). FloraNeotropica Monografia 14: 1–662.

Strehl, T. 1994. Bromelias que passam parte do ano submersas. Bromelia1: 19–21.

Tomlinson, P. B. 1969. Anatomy of the Monocotyledons III. Commelinales-Zingiberales. At The Clarendon Press. Oxford. 446 p.

Varadarajan, G. S. and A. J. Gilmartin. 1987. Foliar scales of the subfamilyPitcairnioideae (Bromeliaceae). Systematic Botany 12: 562–571.

Versieux, L. M. and T. Wendt. 2006. Checklist of Bromeliaceae of MinasGerais, Brazil, with notes on taxonomy and endemism. Selbyana27(2): 107–146.

Versieux, L. M. and T. Wendt. 2007. Bromeliaceae diversity and conser-vation in Minas Gerais state, Brazil. Biodiversity and Conservation 16:2989–3009.

Versieux, L. M., J. V. Coffani-Nunes, G. M. Paggi, and A. F. Costa. Check-list das Bromeliaceae do Mato Grosso do Sul, Brasil. Iheringia. SerieBotanica (in press).

Versieux, L. M., N. Vasconcellos, and G. Martinelli. 2013. Tillandsia bonita(Bromeliaceae), a new species from Mato Grosso do Sul, Brazil, withNotes on Leaf Anatomy. Systematic Botany 38: 75–81.

Versieux, L. M., T. Wendt, R. B. Louzada, and M. G. L. Wanderley.2008. Bromeliaceae da Cadeia do Espinhacoo. Megadiversidade 4:98–110.

Wiesbauer, M. B., E. C. Scariot, L. L. Sasaki, and A. Reis. 2007. Influenciada luz e inundacao na germinacao de Dyckia distachya Hassler,uma bromelia em vias de extincao. Revista Brasileira de Biociencias5(suppl. 1): 717–719.

2015] PAGGI ET AL.: REDISCOVERING DYCKIA EXCELSA 135